Using abundance data to assess the relative role of sampling biases and evolutionary radiations in Upper Muschelkalk ammonoids

The Middle Triassic ammonoid genus Ceratites diversified spectacularly within the Germanic Muschelkalk Basin during the Anisian/Ladian (244–232 Mya). Previous studies have interpreted this diversification as a sequence of rapid, endemic radiations from a few immigrant taxa. Here we investigate the possibility that geological and sampling biases, rather than ecological and evolutionary processes, are responsible for this pattern. A new specimen based dataset of Ceratites species-richness and abundance was assembled. This dataset was combined with 1:200000 geological maps in a geodatabase to facilitate geospatial analyses. One set of analyses compared species richness per geological map with the number of occurrences and localities per map. Per-map change in the amount of rock available to sample for fossils was also included as a variable. Of these three variables, number of occurrences is the most strongly correlated with richness. Variation in the amount of rock is not a strong determinant of species-richness. However, rarefaction of basin-wide species/abundance data demonstrates that differences in species-richness through time are not attributable to sample size differences. The average percent similarity among sites remained close to 50% throughout the Upper Muschelkalk. The rank abundance distribution (RAD) of species from the first interval of the Upper Muschelkalk is consistent with colonization of a disturbed environment, while the other two intervals have RADs consistent with more stable ecosystems. These results indicate that genuine ecological and evolutionary events are partly responsible for the observed differences in richness and abundance. Although changes in the RADs through time support changes in the ammonoid assemblage structure, the processes underlying increasing richness and change in RADS cannot be explained by increasing geographic distinctiveness or isolation among the ammonoid assemblages present at different localities

Reef refugia in the aftermath of past episodes of global warming

In the face of rising global temperatures, coral reefs experience coral mass bleaching and mortality. Subtropical and mesophotic environments may represent refugia for reef corals under climate change, where they can survive and eventually recolonize degraded areas. Using a comprehensive database of fossil reefs, we empirically assess the efficacy of subtropical, deeper, and turbid mesophotic environments to restore coral reefs after past global warming events. We focus on tropical coral reefs over the last 275 million years and four rapid climate warming events, which coincided with global reef crises in the geological record. In the aftermath of such hyperthermal events, we observed an increase in the proportions of reefs occurring in deeper (blue) mesophotic environments. Additionally, we found a trend of reef distributions and coral shifting towards higher latitudes. The number of coral occurrences in turbid (brown mesophotic) environments also increased after hyperthermal events. Our results suggest that subtropical, blue, and brown mesophotic environments may have served as immediate refugia for shallow-water coral species escaping warming seawater. While the patterns of reef range shifts and the establishment of blue and brown mesophotic refugia following ancient hyperthermal events provide some hope for coral reefs under current climate change, re-establishement of background reef conditions took most times millions of years. Ante el incremento de temperatura global, los arrecifes coralinos están experimentando eventos masivos de blanqueamiento y mortalidad. Los ambientes subtropicales y mesofóticos pueden representar refugios para los corales arrecifales, en los cuales pueden escapar de los efectos del cambio climático, sobrevivir y desde allí recolonizar áreas previamente degradadas. Mediante el uso de una exhaustiva base de datos en arrecifes coralinos, en este estudio se evaluó empíricamente la eficacia de los ambientes subtropicales y mesofóticos, tanto de aguas turbias someras (marrones) como de aguas claras profundas (azules), en la recuperación de arrecifes coralinos después de eventos hipertermales en el pasado. Nuestro enfoque estuvo en los arrecifes coralinos tropicales durante los últimos 275 millones de años y cuatro eventos de calentamiento climático rápido, los cuales coinciden con crisis globales en la ocurrencia de arrecifes en el registro fósil. Como consecuencia de dichos eventos hipertermales, observamos un aumento del número de arrecifes en ambientes mesofóticos de aguas profundas (azules). Además, encontramos una tendencia en la distribución de arrecifes y corales que se desplazan hacia latitudes más altas. También se observó un aumento en el número de corales que estuvieron presentes en ambientes de aguas turbias (marrones) después de dichos eventos hipertermales. Nuestros resultados sugieren que, en el pasado, los ambientes subtropicales, mesofóticos azules y mesofóticos marrones pudieron haber servido como refugios inmediatos para las especies de coral de aguas someras, en los cuales encuentran condiciones atenuantes ante el calentamiento oceánico. Si bien los patrones de desplazamiento de los arrecifes en el rango latitudinal y el establecimiento de refugios mesofóticos de aguas marrones y azules posteriores a eventos hipertermales brindan una luz de esperanza para el futuro de los arrecifes coralinos de cara al cambio climático actual, nuestros resultados evidencian que el restablecimiento de los arrecifes puede tomar millones de años

Maximum rates of climate change are systematically underestimated in the geological record

D.B.K. was supported by NERC Fellowship grant NE/I02089X/1 and W.K. by DFG grant Ki 806/12-1. Jonny Beedell is thanked for his help in data compilation and Michael Joachimski for discussions.Peer reviewedPublisher PD

Fast-growing species shape the evolution of reef corals

Ecological interactions are ubiquitous on tropical coral reefs, where sessile organisms coexist in limited space. Within these high-diversity systems, reef-building scleractinian corals form an intricate interaction network. The role of biotic interactions among reef corals is well established on ecological timescales. However, its potential effect on macroevolutionary patterns remains unclear. By analysing the rich fossil record of Scleractinia, we show that reef coral biodiversity experienced marked evolutionary rate shifts in the last 3 million years, possibly driven by biotic interactions. Our models suggest that there was an overwhelming effect of staghorn corals (family Acroporidae) on the fossil diversity trajectories of other coral groups. Staghorn corals showed an unparalleled spike in diversification during the Pleistocene. But surprisingly, their expansion was linked with increases in both extinction and speciation rates in other coral families, driving a nine-fold increase in lineage turnover. These results reveal a double-edged effect of diversity dependency on reef evolution. Given their fast growth, staghorn corals may have increased extinction rates via competitive interactions, while promoting speciation through their role as ecosystem engineers. This suggests that recent widespread human-mediated reductions in staghorn coral cover, may be disrupting the key macroevolutionary processes that established modern coral reef ecosystems

Deep-time climate legacies affect origination rates of marine genera

Biodiversity dynamics are shaped by a complex interplay between current conditions and historic legacy. The interaction of short- and long-term climate change may mask the true relationship of evolutionary responses to climate change if not specifically accounted for. These paleoclimate interactions have been demonstrated for extinction risk and biodiversity change, but their importance for origination dynamics remains untested. Here, we show that origination probability in marine fossil genera is strongly affected by paleoclimate interactions. Overall, origination probability increases by 27.8% [95% CI (27.4%, 28.3%)] when a short-term cooling adds to a long-term cooling trend. This large effect is consistent through time and all studied groups. The mechanisms of the detected effect might be manifold but are likely connected to increased allopatric speciation with eustatic sea level drop caused by sustained global cooling. We tested this potential mechanism through which paleoclimate interactions can act on origination rates by additionally examining a proxy for habitat fragmentation. This proxy, continental fragmentation, has a similar effect on origination rates as paleoclimate interactions, supporting the importance of allopatric speciation through habitat fragmentation in the deep-time fossil record. The identified complex nature of paleoclimate interactions might explain contradictory conclusions on the relationship between temperature and origination in the previous literature. Our results highlight the need to account for complex interactions in evolutionary studies both between and among biotic and abiotic factors

Not all biodiversity rich spots are climate refugia

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Adding fossil occupancy trajectories to the assessment of modern extinction risk

Besides helping to identify species traits that are commonly linked to extinction risk, the fossil record may also be directly relevant for assessing the extinction risk of extant species. Standing geographical distribution or occupancy is a strong predictor of both recent and past extinction risk, but the role of changes in occupancy is less widely assessed. Here we demonstrate, based on the Cenozoic fossil record of marine species, that both occupancy and its temporal trajectory are significant determinants of risk. Based on extinct species we develop a model on the additive and interacting effects of occupancy and its temporal changes on extinction risk. We use this model to predict extinction risk of extant species. The predictions suggest a moderate risk for marine species on average. However, some species seem to be on a long-term decline and potentially at a latent extinction risk, which is not considered in current risk assessments

A possible link between coral reef success, crustose coralline algae and the evolution of herbivory

Crustose coralline red algae (CCA) play a key role in the consolidation of many modern tropical coral reefs. It is unclear, however, if their function as reef consolidators was equally pronounced in the geological past. Using a comprehensive database on ancient reefs, we show a strong correlation between the presence of CCA and the formation of true coral reefs throughout the last 150 Ma. We investigated if repeated breakdowns in the potential capacity of CCA to spur reef development were associated with sea level, ocean temperature, CO2 concentration, CCA species diversity, and/or the evolution of major herbivore groups. Model results show that the correlation between the occurrence of CCA and the development of true coral reefs increased with CCA diversity and cooler ocean temperatures while the diversification of herbivores had a transient negative effect. The evolution of novel herbivore groups compromised the interaction between CCA and true reef growth at least three times in the investigated time interval. These crises have been overcome by morphological adaptations of CCA